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Reproductive Toxicity and Endocrine Disruption Chapter | 17 283
VetBooks.ir and connective tissue (Senger, 2003; Evans and Ganjam, production of pheromone (Haschek et al., 2010). These
accessory sex glands in the male are generally considered
2017). The Leydig cells are homologous to the theca
to be androgen dependent, with conversion of testosterone
interna cells in the ovary and produce testosterone (also
estrogen in some species). There are species differences to DHT occurring in the prostate and seminal vesicles of
with respect to the abundance of Leydig cells in the inter- many species (Senger, 2003; Haschek et al., 2010; Evans
stitium, and these differences are important to recognize and Ganjam, 2017). The weights of the accessory sex
when reporting Leydig or interstitial cell hyperplasia in glands can be used as an indirect measure of testosterone
response to toxicant exposure. It should also be noted that concentrations or exposure to antiandrogens (Thomas and
Leydig and, to a lesser extent, Sertoli cells contain Thomas, 2001; Senger, 2003; Evans and Ganjam, 2017).
enzymes involved in xenobiotic biotransformation, and
the synthesis of toxic metabolites can actually occur External Genitalia
within the testis, in close proximity to the target cells for
The external genitalia of the male consist of the copula-
a given reproductive toxicant (Thomas and Thomas,
tory organ or penis, the prepuce, which protects the penis
2001; Haschek et al., 2010).
from environmental and mechanical injury, and the scro-
tum for testes positioned outside of the abdominal cavity.
Excurrent Duct System Penile structure is extremely species variable, with some
The excurrent duct system consists of the efferent duc- species even having a special penile bone (i.e., os penis),
tules, the epididymal duct and the ductus deferens. This but the shaft of the penis generally consists of erectile tis-
duct system functions to conduct spermatozoa, rete fluid sue (corpus cavernosum and corpus spongiosum) which
and some testicular secretory products away from the tes- surrounds the pelvic urethra. The glans penis is homolo-
tis and eventually into the pelvic urethra (Senger, 2003; gous to the female clitoris, and stimulation of the glans is
Evans and Ganjam, 2017). The reabsorption of fluid by a the primary factor involved in the initiation of ejaculation
species-variable number of efferent ductules is essential (Senger, 2003; Evans and Ganjam, 2017). The scrotum
for normal testicular function (O’Donnell et al., 2001; protects the testes from mechanical injury and, in con-
Hess, 2003), and these tubules terminate by joining a sin- junction with the tunica dartos, cremaster muscle and
gle highly coiled epididymal duct, commonly referred to pampiniform plexus, plays a major thermoregulatory role
as the epididymidis or epididymis. Depending on the spe- with respect to temperature-sensitive, testicular spermato-
cies, the epididymidis is generally subdivided into the ini- genesis. In some species of wildlife (e.g., elephants and
tial segment, head (caput), body (corpus) and tail (cauda), marine mammals), the testes are positioned intra-
with the various portions sometimes being further subdi- abdominally. Xenobiotics, which cause hyperthermia (i.e.,
vided (Franca et al., 2005). The primary functions of the ergopeptine alkaloids) or which induce fever, have the
¸
epididymidis are transport and sustenance of sperm, reab- potential to adversely affect spermatogenesis.
sorption and secretion of fluid (initial segment and head,
respectively); spermatozoal acquisition of motility and Spermatogenesis
fertile potential (i.e., sperm maturation); recognition and
Spermatozoa are highly specialized haploid cells equipped
elimination of defective spermatozoa; sperm storage prior
with a self-powered flagellum to facilitate motility, as
to ejaculation and secretory contributions to the seminal
well as an acrosome to mediate penetration of the zona
fluid (Sutovsky et al., 2001). The epididymal transit time
pellucida. Spermatogenesis takes place within the semi-
varies somewhat with species, but is generally approxi-
niferous tubules and consists of all the changes germ cells
mately 7 to 14 days in length, depending on several fac-
undergo in the seminiferous epithelium in order to pro-
tors including ejaculation frequency. The ductus deferens
duce adequate numbers of viable spermatozoa each day
conducts spermatozoa matured in the epididymidis to the
and to continuously replace spermatogonial stem cells
pelvic urethra which helps to form the penis.
(Thomas and Thomas, 2001; Senger, 2003; Evans and
Ganjam, 2017). Spermatogenesis provides for genetic
Accessory Sex Glands diversity and ensures that germ cells are in an immuno-
There are a number of accessory sex glands (the comple- logically favored site (Senger, 2003; Evans and Ganjam,
ment of which varies with species) that contribute to the 2017). The duration of spermatogenesis varies with spe-
composition of the seminal fluid in mammals. These cies but generally ranges between 4 and 8 weeks (approx-
glands include the ampullae, seminal vesicles (vesicular imately 30 60 days) in domestic and laboratory animals
glands), prostate and bulbourethral glands (Senger, 2003; and is approximately 75 days (almost 11 weeks) in
Evans and Ganjam, 2017). Laboratory rodents (i.e., mice humans. It is important to keep in mind the durations of
and rats) have an additional gland referred to as the pre- spermatogenesis and epididymal sperm transport in a
putial gland, which appears to have a role in the given species, as well as the normal, species-specific